Tormos et al.: Final-instar larva of 333

DESCRIPTION OF THE FINAL INSTAR LARVA OF PERITHOUS SCURRA WITH COMMENTS ON ITS MORPHOLOGICAL CHARACTERS (: ICHNEUMONIDAE, , DELOMERISTINI)

J. TORMOS1, J. D. ASÍS1 AND J. SELFA2 1Unidad de Zoología, Facultad de Biología, Universidad de Salamanca 37071-Salamanca Spain

2Laboratori d’Entomologia i Control de Plagues, Departament de Biologia Universitat de València, Dr. Moliner, 50. 46100 (València) (Spain)

ABSTRACT

A description is given of the final instar-larva of Perithous scurra (Panzer, 1804), which is compared with the previously described species of the genus, P. divinator (Rossius, 1790). The greater or lesser development of the epistoma, and the number of sensilla of the maxillary and labial palpi are the best characters for distinguishing between both species. Additionally, the larva of P. scurra presents spinules on the teg- ument and sensilla on the antennal orbits, but none of those structures has been de- scribed in P. divinator.

Key Words: Ectoparasitoid, final-instar larva, Ichneumonidae

RESUMEN

Se describe la larva madura de Perithous scurra (Panzer, 1804), y se compara con la especie previamente descrita del género, P. divinator (Rossius, 1790). El mayor o menor desarrollo del epistoma, y el número de sensilas de los palpos maxilares y la- biales, son los caracteres que mejor permiten distinguir ambas especies. Adicional- mente, la larva de P. scurra presenta espínulas en el tegumento y sensilas en las órbitas antenales, aunque ninguna de estas estructuras ha sido citada en P. divinator.

The immature instars of parasitic Hymenoptera have been studied by several au- thors, the classic works of Clausen (1940) and Hagen (1964) being outstanding, as are the keys for the taxonomic differentiation of mature larvae compiled by Beirne (1941), Short (1952, 1959, 1970, 1976), Finlayson (1967, 1975) and Capek (1970). Within this broad group, the Ichneumonidae have been particularly studied by Short (1978), Fin- layson & Hagen (1979), Finlayson (1967, 1987), Wahl (1986), and Brooks & Wahl (1987). Here we describe the mature larva of a species of Pimplinae (= Ephialtinae, sensu Townes, 1969) [Delomeristini (sensu Fitton et al., 1988)] [= Theronini auct.]: Perithous scurra (Panzer, 1804) (= mediator Fabricius). This species is a larval ectoparasitoid of Hymenoptera Aculeata that nest in hollow stems (Aubert, 1969), and its biology has been studied by Verhoeff (1891), Bouché (1847), Brocher (1926), and Fitton et al. (1988). Although a detailed description of the internal anatomy of the mature larva was provided by Brocher (1926), he failed to describe the morphological characters present on the cephalic structures (sclerotized areas of the external cephalic skele-

334 Florida Entomologist 82(2) June, 1999 ton), spiracles (usually prothoracic), and tegument, that would permit a characteriza- tion of the preimaginal stages of Hymenoptera Parasitica (Short 1978; Finlayson & Hagen 1979).

MATERIALS AND METHODS

Three mature larvae and six imagos of P. scurra were obtained from two trap nests of Ailanthus altissima Swingle (Simaroubaceae), in which (Shuckard, 1837) (Hymenoptera, Sphecidae) had nested. The trap nests were placed in the field in April 1993 at Casas Valle Carmona (Cuenca, Spain) and were collected in September. The methodology of Asís (1990) was employed to open the nests, and for data collection, preservation, and preparation; the terminology of Finlayson (1987) was used for the description of the mature larva. Voucher specimens are deposited in the Departamento de Zoología, Facultad de Bi- ología, Universidad de Salamanca, Salamanca (Spain).

RESULTS

Perithous scurra (Panzer) Of the eight cells contained in one of the nests of P. lethifer, the externalmost five were parasitized; in the other nest with nine cells the externalmost three and the sixth were parasitized. The mature larva had not constructed a cocoon in any case. One male and two females of the emerged from the first of the nests (two mature larvae were stored in vials with 70% ethanol for later study), while three fe- males emerged from the second one (one larva was preserved for later study). Two of the larvae (ref.: 93070401) were prepared for microscopic examination while the other one (ref.: 93070402 A) was used for whole-body drawings. Mature larva. General aspect (Fig. 1). Elongate (length = 1.25 cm, maximum width = 22 mm), more or less fusiform, slightly curved body, with three thoracic and ten ab- dominal segments; whitish, weakly sclerotized, except spiracles and setae. Anus small, almost terminal. Pleural lobes well developed. Tegument papillose, with spinules (l = 3 mm) and setae (l = 25-40 mm) (Fig. 2); the latter in a transverse line around each of the body segments. Spiracles (Fig. 3) located on prothorax and first eight abdominal segments; atrium (diameter = 30 mm) round, unarmed, separated from closing apparatus (l = 22 mm, w = 15 mm) by a short section of trachea (l = 19 mm, w = 18 mm). Cranium (Fig. 4) (w = 825 mm, h = 630 mm) small, weakly sclerotized, slightly re- tracted within prothorax; with sensilla (d = 6 mm) and sparse setae (l = 13-50 mm), more numerous on lateral side of sclerotized areas of external cephalic skeleton; ecdysial suture line and ocular lines undifferentiated (they are slightly apparent on larva not treated for microscopy); antennae papilliform (l = 32 mm; w =25 mm), with a sensillum (d = 5 mm) on antennal orbit; epistoma completely developed; pleurostoma, hypostoma, lacinial sclerite, hypostomal spur and stipital sclerite well sclerotized; su- perior and inferior pleurostomal processes present; cardo absent; ventral zone of la- bial sclerite twice or three times width of lateral zones; labral sclerite complete, well sclerotized; clypeus with six setae (l = 5-23 mm) and two sensilla (d = 4 mm); labrum with eight setae (l = 18-20 mm) and eighteen sensilla (l = 1-5 mm). Mouthparts (Fig. 5). Mandibles with wide base and blade bifurcate; in this bifur- cated blade, the larger part has teeth on both dorsal and ventral surfaces, the smaller one being unarmed, and with a large posteromedial tooth; maxillae with five setae (l

Tormos et al.: Final-instar larva of Ichneumonidae 335

Figs. 1-3. Mature larva of P. scurra (Panzer). 1. Lateral view of body; 2. Detail of tegument with papilla, spinules and setae; 3. Spiracle (atrium, trachea, spiracular closure).

336 Florida Entomologist 82(2) June, 1999

Figs. 4 & 5. Mature larva of P. scurra (Panzer). 4. Cranium (frontal view); 5. Detail of antenna, sclerotized areas of cranium (the hypostoma has been cut off), clypeus, la- brum and mouthparts.

Tormos et al.: Final-instar larva of Ichneumonidae 337

= 3-20 mm); labium with four setae (l = 3-20 mm); maxillary (w = 15 mm) and labial (w = 16 mm) palpi not protruding, disc shaped with four sensilla on each; salivary orifice transverse ovoid; spinneret undifferentiated; postlabium with four setae (l = 20 mm).

DISCUSSION The results obtained in the present work confirm the data reported by Wolf (1953) and Fitton et al. (1988) concerning the hosts and the parasitic behaviour of P. scurra. According to Fitton et al. (1988), at least the British species of the genus Perithous Holmgren, 1859 would have become specialist idiobiont ectoparasitoids of sphecids, parasitizing their prepupal stage. In Great Britain, P. scurra parasitizes (Fabricius, 1793), pupating inside the cell of its host without constructing a cocoon. Although these authors consider P. lugubris as a usual host of P. scurra, Wolf (1953) obtained this pimpline from P. lethifer. As in the case of the larvae of the Pimplinae (Short 1978; Finlayson 1967, 1975, 1987), the mature larva of P. scurra has the hypostomal spur joining the stipital scler- ite very close to where the latter joins the labial sclerite. As in the other Delomeristini, the characters described by Short (1978) are well defined; and as an ectoparasitoid species the mandible blades have teeth and a broad unarmed projection in the form of a tooth on the base and blade of the mandibles, nine pairs of functional spiracles (one prothoracic and eight abdominal), and an undifferentiated spinneret. Unlike the previously described species of the genus, P. divinator (Danks 1970; Short 1978), P. scurra displays the following traits (Table 1): a) more setae on clypeus, maxilla, and postlabium; and more sensilla on clypeus and labrum; b) completely de- veloped epistoma [a character only present, among the Delomeristini, in Pseudo- rhyssa Merrill, 1915 (Short 1978)]; c) maxillary and labial palpi with four sensilla (character present in only this species among Delomeristini); d) undifferentiated spin- neret [in P. divinator, although not very differentiated, the spinneret is perceptible]; e) tegument with spinules [this character was not reported either by Danks (1970) or Short (1978) for P. divinator]; and f) antennal orbit with a single, well differentiated sensillum. Furthermore, in the species studied the pleurostoma is well sclerotized, as in P. divinator and Pseudorhyssa, and there is no seta on the posterior part of the stipital sclerite, the cardo being absent. These latter two characters were reported by Short (1978) for P. divinator, although Danks (1970) did not include them in his description. In any case, it should be noted that the comparison has been made with Short’s (1970) and Dank’s (1978) descriptions of P. divinator; according to Brooks & Wahl (1987), those descriptions could be inadequate in several aspects.

TABLE 1. MORPHOLOGICAL DIFFERENCES BETWEEN THE MATURE LARVAE OF P. SCURRA (PANZER) AND P. DIVINATOR (ROSSI). CHARACTERS [PRESENCE (*); ABSENCE (-)]: NUMBER OF SETAE ON: 1) CLYPEUS; 2) MAXILLAE; 3) POSTLABIUM; NUM- BER OF SENSILLA ON: 4) CLYPEUS; 5) LABRUM; 6) MAXILLARY AND LABIAL PALPI; 7) EPISTOMA COMPLETELY DEVELOPED; 8) SPINNERET; 9) SPINULES ON TEGUMENT; 10) SENSILLUM OF ANTENNAL ORBIT.

Species 12345678910

P. scurra 6646184*—** P. divinator 4 3/4 2 4 2 3 — * ? ?

338 Florida Entomologist 82(2) June, 1999

ACKNOWLEDGMENTS We are much indebted to Thelma Finlayson (Simon Fraser University, Burnaby, Canada), for her comments on the manuscript; David Wahl provided also some help in the final version of the draft. Grants from de Junta de Castilla y León (SA 18/96) supported the study.

REFERENCES CITED

ASÍS, J. D. 1990. Biología de esfécidos ibéricos (Hymenoptera: Sphecidae). Tesis Doc- toral. Universitat de València (inédita). AUBERT, J. F. 1969. Les ichneumonides ouest-Paléarctiques et leurs hôtes. 1. Pimpli- nae, Xoridinae, Acaenitinae. Quatre Feuilles Editeur. France. BEIRNE, B. P. 1941. A consideration of the cephalic structures and spiracles of the fi- nal instar larvae of the Ichneumonidae (Hym.) Trans. Soc. Br. Ent. 7: 123-190. BOUCHÉ, P. F. 1847. Beiträge zur Kenntnis der Insekten-Larven. Ent. Ztg. Stettin 8: 162-165. BROCHER, F. R. 1926. Observations sur la Perithous mediator Gr. Ponte, Oeuf, larve. Nymphe et Imago. Etude anatomique de la tarière, de ses muscles et de son fonctionnement. Ann. soc. ent. France 95: 393-410. BROOKS, R. W., AND D. B. WAHL. 1987. Biology and mature larva of Hemipimpla pul- chripennis (Saussure), a parasite of Ropalidia (Hymenoptera: Ichneumonidae, Vespidae). J. New York Entomol. Soc. 95(4): 547-552. CAPEK, M. 1970. A new classification of the Braconidae (Hymenoptera) based on the cephalic structures of the final instar larva and biological evidence. Can. Ent. 102: 846-875. CLAUSEN, C. P. 1940. Entomophagous . McGraw-Hill. New York and London. DANKS, H. V. 1970. Biology of some stem-nesting aculeata Hymenoptera. Trans. R. Entomol. Soc. London 122: 323-399. FINLAYSON, T. 1967. A classification of the subfamily Pimplinae (Hymenoptera: Ich- neumonidae) based on final-instar larval characteristics. Can. Ent. 99: 1-8. FINLAYSON, T. 1975. The cephalic structures and spiracles of final-instar larvae of the Subfamily , Tribe Campoplegini (Hymenoptera: Ichneu- monidae). Mem. ent. Soc. Can. 94: 1-137. FINLAYSON, T. 1987. , pp. 602-617, 649-665. in F. W. Stehr (ed.) Im- mature insects. Kendall/Hunt Publishing Company, Dubuque, Iowa. FINLAYSON, T., AND K. S. HAGEN. 1979. Final-instar larvae of parasitic Hymenoptera. Pest Management Paper N° 10. Department of Biological Sciences, Simon Fraser University, Burnaby, B.C. FITTON, M. G., M. R. SHAW, AND I. D. GAULD. 1988. Pimpline ichneumon-flies. Hy- menoptera, Ichneumonidae (Pimplinae). Handbooks for the Identification of British Insects 7 (1): 1-110. HAGEN, K. S. 1964. Developmental stages of parasites, pp. 186-246 in P. Debach (ed.) Biological Control of pests and weeds. Chapman and Hall. London. SHORT, J. R. T. 1952. The morphology of the head of larval Hymenoptera with special reference to the head of Ichneumonoidea, including a classification to the final instar larvae of the Braconidae. Trans. R. Entomol. Soc. Lond. 103: 27-84. SHORT, J. R. T. 1959. A description and classification of the final instar larvae of the Ichneumonidae (Insecta, Hymenoptera). Proc. U.S. Natn. Mus. 110 (3419): 391- 511. SHORT, J. R. T. 1970. On the classification of the final instar larvae of the Ichneu- monidae (Hymenoptera). Supplement. Trans. R. Entomol. Soc. London 112: 185-210. SHORT, J. R. T. 1976. A description and classification of some final-instar larvae of the Mesochorinae (Hymenoptera, Ichneumonidae). Syst. Entomol. 1: 195-200. SHORT, J. R. T. 1978. The final larval instars of the Ichneumonidae. Mem. American Entomol. Inst. 25. 508 pp. Shelly: Aggression in Female Fruit Flies 339

TOWNES, H. 1969. The genera of Ichneumonidae, part 1. Memoirs of the American En- tomological Institute 11: 1-300. VERHOEFF, C. 1891. Biologische Aphorismen über einige Hymenopteren, Dipteren und Coleopteren. Verh. naturh. Ver. Reinl. 48: 1-80. WAHL, D. B. 1986. Larval structures of oxytorines and their significance for the higher classification of some Ichneumonidae (Hymenoptera). Syst. Entomol. 117-127. WOLF, H. 1953. Beiträge zur Hymenopterenfauna des oberen Lahn-Dill-Sieg-Gebites. Nachr. naturw. Mus. Aschaffenburg 41: 83-85.

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DEFENSE OF OVIPOSITION SITES BY FEMALE ORIENTAL FRUIT FLIES (DIPTERA: TEPHRITIDAE)

TODD E. SHELLY1 Hawaiian Evolutionary Biology program, University of Hawaii Honolulu, Hawaii, 96822

1Current address: Organization for Tropical Studies, Duke University 410 Swift Avenue, Durham, NC 27705

ABSTRACT

Field observations revealed that females of the oriental fruit fly, Bactrocera dorsa- lis (Hendel), defended oviposition sites on mangos (Mangifera indica L.) against con- specific females. In most encounters, females simply lunged at opponents and chased them off the fruit without physical contact. However, head-butting and pushing were observed in about 10% of the contests. Body size was a key determinant of fighting success, with larger females winning 85% of the encounters. In a field experiment, ar- rivals, oviposition, and aggression of females were compared between intact vs. sliced peaches. Similar numbers of females landed on the two classes of fruits, but a greater proportion of alighting females oviposited on sliced peaches than intact peaches. The adaptive function of female territoriality is discussed in light of these findings.

Key Words: fruit fly, aggression, territory, oviposition, Hawaii

RESUMEN Observaciones de campo han indicado que las hembras de la mosca oriental de la fruta, Bactrocera dorsalis (Hendel), defienden los sitios de oviposición en mangos (Mangifera indica L.) de las hembras de su misma especie. En la mayoría de los en- cuentros, las hembras simplemente se lanzaron hacia sus oponentes y las espantaron de la fruta sin tener contacto físico directo. Sin embargo, choques de cabeza y empu- jones fueron observados en el 10% de los encuentros. El tamaño del cuerpo es clave en la determinación del éxito en las luchas; éste se refleja en que las hembras más gran- des ganaron en un 85% de los encuentros. En un experimento de campo, llegadas, ovi-